new omomyoids (euprimates, mammalia) from the late uintan

Palaeontologia Electronica palaeo-electronica.org

New omomyoids (Euprimates, Mammalia) from the late Uintan of southern California, USA, and the question of

the extinction of the Paromomyidae (Plesiadapiformes, Primates)

Sergi López-Torres, Mary T. Silcox, and Patricia A. Holroyd

ABSTRACT

Paromomyidae has been thought to represent the longest-lived group of stem pri-mates (plesiadapiforms), extending from the early Paleocene to late Eocene. We ana-lyzed primate material from the late-middle Eocene of southern California that hadinitially been ascribed to cf. Phenacolemur shifrae. This material falls at the lowest endof the size range for the family. The Californian specimens also exhibit several dentalfeatures that are atypical for paromomyids, such as a strong paraconid on the thirdlower molar, and differ from early Eocene species of Phenacolemur in lacking a distallyexpanded distolingual basin on upper molars. This combination of traits is more typicalof earlier paromomyids with more plesiomorphic morphologies (e.g., Paromomys) andas such is inconsistent with the late age of these specimens. The purported paromo-myids Ph. shifrae and Ignacius mcgrewi known in deposits of similar age are compara-bly tiny and share many of the characteristics found in the southern California materialthat distinguish them from typical early Eocene paromomyids. However, these traitsare shared with some trogolemurin omomyoid euprimates that are of similar size. Weargue that the material from southern California, along with Ph. shifrae and I. mcgrewi,should be transferred to a new genus of trogolemurin omomyoid. Purported Europeanrecords of paromomyids later than the earliest middle Eocene are reconsidered andfound to be non-diagnostic. After the early middle Eocene, only a single tooth of aparomomyid can be confirmed, indicating that the group suffered near-extinction, pos-sibly correlated with the Early Eocene Climatic Optimum.

Sergi López-Torres. University of Toronto Scarborough, Department of Anthropology, 1265 Military Trail, Toronto, ON, M1C 1A4, Canada. Current address: Polish Academy of Sciences, Roman Kozłowski Institute of Paleobiology, Department of Evolutionary Paleobiology, Twarda 51/55, Warsaw 00-818, Poland. [email protected] T. Silcox. University of Toronto Scarborough, Department of Anthropology, 1265 Military Trail, Toronto, ON, M1C 1A4, Canada. [email protected]

http://zoobank.org/5EE56527-5F98-4724-8120-18F7471BB497

López-Torres, Sergi, Silcox, Mary T., and Holroyd, Patricia A. 2018. New omomyoids (Euprimates, Mammalia) from the late Uintan of southern California, USA, and the question of the extinction of the Paromomyidae (Plesiadapiformes, Primates). Palaeontologia Electronica 21.3.37A 1-28. https://doi.org/10.26879/756palaeo-electronica.org/content/2018/2309-new-primate-from-california

Copyright: September 2018 Society of Vertebrate Paleontology. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.creativecommons.org/licenses/by/4.0/creativecommons.org/licenses/by-nc-sa/4.0/

LÓPEZ-TORRES, SILCOX, & HOLROYD: NEW PRIMATE FROM CALIFORNIA

Patricia A. Holroyd. University of California Museum of Paleontology, 1101 Valley Life Sciences Building, Berkeley, CA, 94720, USA. [email protected]

Keywords: Omomyoidea; Trogolemurini; Paromomyidae; Eocene; new genus; new species

Submission: 21 January 2017 Acceptance: 23 August 2018

INTRODUCTION

The Paromomyidae is a family of extinct fossilmammals known from the early Paleocene (Torre-jonian 1) to the late Eocene (late early Chadronian)of North America (Silcox and Gunnell, 2008; Silcoxet al., 2008; Clemens and Wilson, 2009; Kihm andTornow, 2014), the early through middle Eocene ofEurope (Russell et al., 1967; Godinot, 1984;Estravís, 2000; Aumont, 2003; Hooker, 2010;Marigó et al., 2012, 2014), and the early Eocene ofAsia (Tong and Wang, 1998). Paromomyids havebeen characterized dentally as possessingenlarged lower incisors that are inclined nearly-hor-izontally (i.e., sub-horizontal); P4 with a tall,pointed, broad-based protoconid; low crownedmolars; and a large hypoconulid lobe on M3 (Silcoxand Gunnell, 2008, Silcox et al., 2017; e.g., Phena-colemur citatus in Figure 1). Cranially, paromomy-ids have a long snout, small and widely spacedorbits, absence of postorbital bar, wide zygomaticarches, and inflated auditory bullae ossified fromthe entotympanic (Silcox and Gunnell, 2008). Thepostcranial skeleton of Paromomyidae suggestsarborealism and vertical climbing, with stronggrasping abilities similar to those of Ptilocercus, butno gliding or suspensory behaviors (Bloch andBoyer, 2007; Boyer and Bloch, 2008). FollowingSilcox et al. (2017), seven genera comprise thefamily: Paromomys, Phenacolemur, Ignacius,Elwynella, Arcius, Acidomomys, and Edworthia.

In a phylogenetic context, paromomyids werefound to be closely related to Picrodontidae andthe polyphyletic assemblage “Palaechthonidae”, allbeing placed together under the superfamily Paro-momyoidea (Silcox, 2001; Silcox and Gunnell,2008; Silcox et al., 2017). Whereas, some authorshave suggested that paromomyids are moreclosely related to dermopterans than to primates(Beard, 1989, 1990, 1993a, 1993b; Kay et al.,1990, 1992; Ni et al., 2013), other comprehensiveanalyses have provided compelling evidence thatthey are instead stem primates (Krause, 1991;Runestad and Ruff, 1995; Stafford and Thorington,1998; Hamrick et al., 1999; Silcox, 2001; Sargis,

2002; Bloch and Boyer, 2007; Bloch et al., 2007;Boyer and Bloch, 2008).

Paromomyids are extensively distributedacross North America, with records from as farnorth as Ellesmere Island, and as far south asTexas (Schiebout, 1974; West and Dawson, 1977;McKenna, 1980; Eberle and Greenwood, 2012).Among the furthest westward proposed NorthAmerican records of the family is materialdescribed by Mason (1988, 1990) and Walsh(1991a) from Ventura and San Diego counties(California) that was referred to Phenacolemur andcompared to the species Ph. shifrae Krishtalka1978, previously identified from late middle Eocenedeposits in Wyoming assigned to the Uintan and

FIGURE 1. The late Wasatchian paromomyid Phena-colemur citatus. 1: USGS 6573 (original fossil), rightmaxilla with P3-M2. 2-4: USGS 21712 (cast), left den-tary with P4-M3 in occlusal (2), buccal (3), and lingual

(4) views.

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Duchesnean North American Land Mammals Ages(see Robinson et al., 2004).

However, it is worth noting that there havebeen some instances of confusion in late occurringsamples between paromomyids and another groupof early Cenozoic primates, the Omomyoidea.Omomyoids are euprimates (i.e., primates of mod-ern aspect, more closely related to living primatesthan to plesiadapiforms) that first appear near thePaleocene-Eocene boundary in North America,Europe, and Asia (Ni et al., 2004; Smith et al.,2006; Rose et al., 2011). During the early Paleo-gene, paromomyid plesiadapiforms and omo-myoid primates both diversified at small bodysizes, and selected subsets of each clade exhibitvery similar dental features (i.e., expanded distolin-gual basins in upper molars, enlarged sub-horizon-tal first lower incisors, large hypoconulid lobes onM3) and may have had similar ecological roles. Alate middle Eocene primate from Saskatchewanoriginally described as a paromomyid (Storer,1990) was found to be a trogolemurin omomyid,Trogolemur leonardi (Beard et al., 1992). Here wefollow a restricted concept of Trogolemurini (i.e.,only containing two genera: Trogolemur andSphacorhysis) as defined by Gunnell and Rose(2002). As so defined, Trogolemurini is a tribe ofanaptomorphine omomyoids known from the lateearly Eocene (late Wasatchian) of Wyoming (Wil-liams and Covert, 1994) and the middle Eocene(Bridgerian to Duchesnean) of Wyoming (Matthew,1909; Beard et al., 1992; Gunnell, 1995), Nevada(Emry, 1990), and Saskatchewan (Storer, 1990).Trogolemurins are among the smallest knownomomyoids. As is true of paromomyid plesiadapi-forms, trogolemurins are characterized by havingenlarged sub-horizontal central incisors, althoughthey sometimes retain a small I2. Consequently,they have an anteriorly deep mandible to accom-modate this enlarged tooth, like paromomyids do.They retain a small canine and a P3, unlike mostparomomyids. The teeth between I1 and P4 arevery close together and are inclined mesially. Thetrogolemurin P4 is reduced in size, with a verysmall to absent metaconid, and the M3 has a shortand broad hypoconulid lobe (Gunnell and Rose,2002; Figure 2), making these teeth quite differentfrom those of most paromomyids, in which the P4

is expanded and the M3 hypoconulid more

enlarged. The P4 paracone is much taller than itsvery small protocone (Gunnell, 1995). Uppermolars of trogolemurins have prominent proto-cones, narrow postprotocingula, and weak conules

(Gunnell and Rose, 2002), making them moreparomomyid-like and generating potential confu-sion when isolated teeth are found (e.g., Storer,1990). No trogolemurin postcranial material hasbeen reported.

The first attempt to place a trogolemurin in acladistic context was done by Williams (1994), andshe found that Tr. myodes was most closely relatedto Anemorhysis. That was also supported by Wil-liams and Covert (1994). Gunnell (1995) depicteda Tetonoides-Arapahovius clade as sister group toTrogolemurini. Although this differs from the resultsof Williams (1994) and Williams and Covert (1994),this conclusion would still ally trogolemurins withNorth American anaptomorphins. Later, larger andmore comprehensive analyses suggested that Tr.myodes is closely related to the European micro-choerines Microchoerus, Necrolemur, Nannopi-thex, and Pseudoloris (Ross et al., 1998; Ni et al.,2004). However, the analysis by Tornow (2008)

FIGURE 2. Holotype of Trogolemur myodes, AMNH12599 (Matthew, 1909: plate LII, figure 5). Right dentarywith P2-M3. 1: occlusal view; 2: buccal view; 3: lingual

view.

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provided support for the previous idea of Trogole-mur myodes grouping with Anemorhysis. Thereare, therefore, two conflicting interpretations of tro-golemurin relationships: they are either related toNorth American anaptomorphins (Williams, 1994;Williams and Covert, 1994; Gunnell, 1995; Tornow,2008) or to European microchoerines (Ross et al.,1998; Ni et al., 2004).

In light of the past confusion between speci-mens of paromomyids and trogolemurins, we for-mally describe and re-examine the affinities of thesouthern Californian primate specimens that hadbeen attributed to cf. Ph. shifrae, and of similarlyaged material from the Uintan and Duchesnean ofWyoming (Robinson, 1968; Krishtalka, 1978).Because these represent some of the youngestreported paromomyids, our study also promptsreconsideration of the larger question of the tempo-ral distribution of paromomyids and, more gener-ally, of the ecological and environmental factorsthat might have been driving these patterns.

Institutional Abbreviations

AMNH–American Museum of Natural History, NewYork City, NY, USA; CM–Carnegie Museum of Nat-ural History, Pittsburgh, PA, USA; LACM–NaturalHistory Museum of Los Angeles County, Los Ange-les, CA, USA; LACM (CIT)–California Institute ofTechnology collections, now held by LACM; RSM–Royal Saskatchewan Museum, Regina, SK, Can-ada; SDSNH–San Diego Society of Natural His-tory, San Diego, CA, USA; SMNH–SaskatchewanMuseum of Natural History (now RSM); UCM–Uni-versity of Colorado Museum, Boulder, CO, USA;UCMP–University of California Museum of Paleon-tology, Berkeley, CA, USA; UM–University of Mich-igan, Ann Arbor, MI, USA; USNM–United StatesNational Museum, Washington, DC, USA; YPMVP–Division of Vertebrate Paleontology, Yale Pea-body Museum, New Haven, CT, USA.

Historical and Geological Context of Southern Californian Localities

The history of collecting fossil mammals fromthe Eocene of southern California dates back tothe work of Stock (1934a, 1934b, 1935, 1936,1938, 1948) and Wilson (1935a, 1935b, 1940a,1940b), when they described the first Eoceneinsectivores, marsupials, and rodents of the area.Later work recognized several local faunas (Lind-say, 1968; Golz, 1976; Golz and Lillegraven, 1977;Lillegraven, 1979, 1980; Eaton, 1982; Kelly, 1990;Mason, 1988, 1990; Kelly et al., 1991; Theodor,1999; Wesley and Flynn, 2003; Ludtke and Proth-

ero, 2004; Colbert, 2006; Tomiya, 2013). Of partic-ular note in the current context are the extensivecontributions by Walsh (Walsh and Estes, 1985;Walsh, 1987, 1991a, 1991b, 1996, 1997, 1998,2000; Walsh and Gutzler, 1999), who amassedimpressive collections that currently reside at theSan Diego Natural History Museum, much of whichremains unpublished. However, the primate mate-rial from southern California is remarkably scarce.The current study deals specifically with materialoriginally collected by Mason (1988, 1990) andWalsh (1991a) from Ventura and San Diego Coun-ties (California). In the Eocene, fossil localities fromthese two Californian counties would have hadsimilar latitudinal positions to today but would havelain approximately 2 degrees of latitude to thesouth and 3 degrees of longitude to the east (seepaleoposition reconstruction in Figure 3) due tomovement along faults. The western US coastlinewould have been approximately 10 degrees to theeast, having since moved west due to post-Eocenecontinental and sea-floor spreading movements(see McQuarrie and Wernicke, 2005 for a recentsynthesis).

FIGURE 3. Reconstruction of western North Americafrom 40 million years ago. Orange dot indicates thepaleoposition of the sites discussed in this paper,whereas the red dot indicates the current relative posi-tion of the sites. The map is reproduced with R. Blakey’spermission (Colorado Plateau Geosystems, Inc.).

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Here we examine specimens from five locali-ties (SDSNH localities 3276, 3426, 4081, 4082,and LACM (CIT) 180), which occur in river-depos-ited (alluvial) terrestrial sediments. These sedi-ments formed on a coastal plain that would havefeatured a mix of lagoonal, estuarine, and riverineenvironments. Westward, the area grades intoshelf sediments deposited in forearc basins cre-ated by the subduction of the Pacific plate underthe North American plate (e.g., Link et al., 1979;Howard and Lowry, 1995; Abbott, 1999; Berry,1999). The shoreline was never very far to thewest, with uplands and mountains with mixedconiferous forests to the east (Frederiksen, 1991).In the Eocene, the climate would have been tropi-cal to subtropical, although researchers differ intheir interpretations of forest structure, seasonality,and amount of precipitation in the various forma-tions that have produced mammalian fossils, asdiscussed below.

The geologic history of San Diego and Ven-tura counties in southern California is complex,having been affected by significant movementalong the San Andreas and related faults, and theliterature can be confusing, as some formationnames as well as age interpretations havechanged. Here we summarize the geology and col-lecting history of five localities that have producedprimate fossil specimens (SDSNH localities 3276,3426, 4081, 4082, and LACM (CIT) 180), providedata relevant to determining their age and paleoen-vironment, and refer the reader to other reviewsthat can provide more in-depth background onalternate interpretations or older data.Mission Valley Formation, southwest SanDiego County (SDSNH localities 3426 and4020). SDSNH locality 3426, Collwood South, is a2 m thick light brown sandy mudstone bed thatoccurs within the restricted concept of the MissionValley Formation (see discussion in Walsh et al.,1996). According to the SDSNH records, ~1150 kgof sediment were processed for microvertebrates.The fauna of SDSNH locality 3426 is united withthat of other nearby localities as the Cloud 9 localfauna, and is considered late Uintan in age (Walsh,1991a, 1996).

Walsh et al. (1996) reported a single-crystal40Ar/39Ar age of 42.83 ± 0.24 Ma from a pink ben-tonite of normal magnetism in the Mission ValleyFormation. Measured sections of the Mission Val-ley Formation in San Diego span both normal andreversed polarity intervals (Bottjer et al., 1991;Walsh et al., 1996). Bottjer et al. (1991) interpretedit as Chron 18r and thought that Chron 19r was

likely missing in the local area (or represented byan unsampled formation), but additional samplingand correlation of local sections led to interpreta-tion as Chrons 20n and 19r (Walsh et al., 1996;Prothero, 2001a), an assignment consistent withthe radiometric age.

Peterson and Abbott (1979) reviewed the geo-logic evidence for immaturity in the development ofclay minerals combined with the common develop-ment of caliche horizons in the Mission Valley For-mation and the slightly older Friars Formation tosuggest that the climate was warm (18-20 ºC meanannual temperature) with low average annual rain-fall (ca. 63 cm3). Additional lines of evidenceinclude pollen documenting the presence of palmsand a diversity of paratropical to tropical tree taxa(Frederiksen, 1991). Additionally, an analysis of themammalian diversity (Novacek and Lillegraven,1979) suggested similarities to complex, moderneastern African habitats with a mix of riparian habi-tats grading into gallery forests and/or savanna, aslightly wider range of warm temperatures (20-22ºC), and possible annual rainfall of 50-100 cm3.Based on land snails from Mission Valley and Fri-ars formations in San Diego, and locality LACM(CIT) 180 in the Sespe Formation, Roth (1988)inferred the presence of dry tropical forest through-out the region.

SDSNH locality 4020 (SR 125 North [Unit I]Grossmont Summit) has previously been assignedto the late Uintan Cloud 9 local fauna (Walsh,1996), and Penkrot and Zack (2016) have reportedon erinaceomorph lipotyphlan tarsals from this site.SDSNH 4020 is in the Mission Valley Formation inthe city of El Cajon. Fossils occurred in a ~0.6-1.5m thick massive, brown, medium-grained, siltysandstone with some calcareous concretions fromwhich thousands of fossils, including articulatedskeletons, were recovered. The presence of pul-monate (terrestrial) snails led the site to be infor-mally called “the snail beds”. Approximately 3265kg of sediment were screenwashed.Member C of Santiago Formation, northwestSan Diego County (SDSNH localities 3276,4081, 4082, and 4925). The only paleoenviron-mental interpretation based exclusively on speci-mens from the Santiago Formation is a study ofland snails from SDSNH locality 3276 (Jeff’s Dis-covery, Santiago Formation, Member C, Oceans-ide, San Diego County, CA), which found thedistribution of shell sizes and shapes to be consis-tent with interpretations of subtropical to tropicalconditions and paleotemperatures in excess of 25ºC (Roth, 1991).

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Prothero (2001b) found the strata of MemberC of the Santiago Formation at the Jeff’s Discoverylocality to be entirely reversed and suggested itcorrelated with Chron C19r (42.5 Ma-41.5 Ma),because correlative rocks with similar fossils fromthe upper Mission Valley Formation in southernSan Diego County probably also correlate with thismagnetic chron (Walsh et al., 1996).

SDSNH localities 4081 and 4082 (EmeraldRidge Sites 1 and 2) and 4925 (Kelly Ranch Core –Mammal and Coprolite Site) have not previouslybeen reported in the literature or assigned to alocal fauna. All are from Member C of the SantiagoFormation and were found during the course ofconstruction projects for the named housing devel-opments in the city of Carlsbad. SDNSH 4081(Emerald Ridge Site 1) is a 0.3-0.6 m thick weakly-cemented sandstone channel with mudstone andsiltstone clasts. A total of ~1995 kg of matrix werecollected and washed through 24 mesh (0.7 mm)screens (SDSNH records). SDSNH 4082 (EmeraldRidge Site 2) is lithologically similar to SDSNH4081 and may be the same bed but is separatedfrom it by faulting. A total of ~1815 kg of matrixwere collected and processed (SDSNH records).SDSNH 4925 is a ~0.6 m thick, gray-green, finegrained sandy siltstone from which approximately~2900 kg of sediment were hand quarried andscreenwashed (SDNSH records).Sespe Formation, Simi Valley, Ventura County(LACM (CIT) 180). Kelly (1990) placed the taxafrom the Dry Canyon localities, including LACM(CIT) 180, as part of the Tapo Canyon local fauna,where it co-occurs with the omomyoid Dyseolemurpacificus in the late Uintan part of the middle mem-ber of the Sespe Formation, the oldest part of themiddle Eocene sequences in the Simi Valley area.Locality LACM (CIT) 180 is situated 389 m abovethe base of the Sespe Formation and wasassigned a late Uintan age (Mason, 1990).

Prothero et al. (1996) placed these rocks inthe lower half of Chron 18r, but Walsh (1996)expressed skepticism about this younger ageassignment. Based on biostratigraphy, Robinson etal. (2004) assigned the Tapo Canyon local fauna toUi3. Most recently, Kelly et al. (2012) proposed acorrelation to Chron 19r based on a new interpreta-tion of the correlation between the southern Cali-fornia middle Eocene record and that of the Uintaand Duchesne River formations in Utah, and thiscorrelation was also found to be most consistentwith the ranges of carnivoramorphs (Tomiya,2013).

As inferred for the Mission Valley Formation,Roth (1988) used the record of land snails andconcluded that dry tropical forest was present inthe Sespe Formation.

In sum, biostratigraphic, magnetostrati-graphic, and radiometric evidence most stronglysuggest that all these southern Californian sitesare similar in age, likely assignable to the Ui3 sub-zone of the Uintan North American Land MammalAge, fall within Chron 19r, i.e., are 42.301 Ma-41.390 Ma in age (Ogg, 2012), and are within thelatest part of the Lutetian Standard Stage. Alterna-tive interpretations that suggest the VenturaCounty localities are younger, or that all the sitesare older (see Tomiya, 2013 for a review), are notwell supported.

SYSTEMATIC PALEONTOLOGY

Order PRIMATES Linnaeus, 1758Suborder HAPLORHINI Pocock, 1918

Superfamily OMOMYOIDEA Trouessart, 1879Family OMOMYIDAE Trouessart, 1879

Subfamily ANAPTOMORPHINAE Cope, 1883Tribe TROGOLEMURINI Szalay, 1976

Genus WALSHINA gen. nov.Figures 4.1-4.4, 5.1-5.11, 6.1-6.19

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1968 Phenacolemur (Matthew, 1915); Robinson, p.324

1976 Ignacius (Matthew and Granger, 1921); Bownand Rose, p. 112 (in part)

1978 Phenacolemur (Matthew, 1915); Krishtalka, p.338, fig. 2-4

1990 Phenacolemur (Matthew, 1915); Mason, p. 2,fig. 2

1991a cf. Phenacolemur (Matthew, 1915); Walsh, p.166, table 1

1996 Phenacolemur (Matthew, 1915); Walsh, p. 85,table 2

Type species. Walshina esmaraldensis gen. et sp.nov.Included species. Type species, Walshinamcgrewi comb. nov. (= I. mcgrewi Robinson,1968), and Walshina shifrae comb. nov. (= Ph.shifrae Krishtalka, 1978).Distribution. Uintan and Duchesnean of Wyo-ming, and Uintan of California.Etymology. In memory of Stephen L. Walsh of theSan Diego Museum of Natural History, in recogni-tion of his work on the San Diego County faunas.Diagnosis. Paracristid of M1 relatively long as inTrogolemur and Sphacorhysis, but paraconid lessclearly distinct from the paracristid. Differs from

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Sphacorhysis (but not Trogolemur) in having lowermolar talonid basins that are relatively deep withsmooth enamel. Differs from Trogolemur (but notSphacorhysis) in that the cristid obliqua of M1 con-tacts the postvallid distal to the protoconid ratherthan between the protoconid and metaconid.Unlike the other trogolemurins, M1 and M2 ofWalshina gen. nov. have strong hypoconulids withdistinct foveae located below and buccal to thehypoconulid. As in Sphacorhysis, the distal aspectof M1 and M2 is convex, whereas in Trogolemur itis concave. M3 hypoconulid narrower than in othertrogolemurins. M3 trigonid significantly taller than inSphacorhysis (but not Trogolemur). Like Sphaco-rhysis, lower molar entocristids form a roundedcontour (i.e., forming a U-shaped entocristid) in lin-gual view, in contrast to the V-shaped entocristid inTrogolemur. Further differs from Trogolemur andSphacorhysis in having much weaker buccal cin-gulids. Notably stronger precingulum on M2 than inTrogolemur. Protocone lingual expansion on theupper molars not as pronounced as in Trogolemur.Compared to Trogolemur, mesial aspect of M3

straighter, and the lingual border of that tooth muchshorter mesiodistally relative to its buccal border.

Discussion. All trogolemurins share a distallyexpanded distolingual basin of the upper molars(particularly marked in W. mcgrewi comb. nov.),which is quite similar to that observed in paromo-myid plesiadapiforms. This similarity is likely onereason why some members of Walshina gen. nov.have previously been considered paromomyids.However, in other ways the morphology of trogole-murins is inconsistent with that of paromomyids,including the presence of paraconids on M3.Walshina gen. nov. remains quite poorly known,with the only record being isolated upper and lowermolars. One likely reason for this limited record isthat the genus includes the smallest North Ameri-can omomyoids (see below).

Walshina esmaraldensis gen. et sp. nov.Figures 4.1-4.4, 5.2, 5.5, 6.1-6.19

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Phenacolemur cf. shifrae Mason, 1990; Walsh,1996.Holotype. LACM 40198, left M1.Hypodigm. M2: SDSNH 42268, 62850, 87336,87337; M3: SDSNH 76267; M1: SDSNH 76337,87331; M2: SDSNH 76338, 87332, 87333; M3:SDSNH 72583, 76266, 76339, 87334, 87335.Type locality. LACM (CIT) 180, north of Simi Val-ley, Ventura County, California, USA. Late Uintan(Ui3). Etymology. In reference to the localities of Emer-ald Ridge (Vulgar Latin: esmaraldus).Diagnosis. Intermediate in size between W.mcgrewi comb. nov. and W. shifrae comb. nov. Dif-fers from W. mcgrewi comb. nov. in having a hypo-conulid closer to the hypoconid than to theentoconid on M2, similar to W. shifrae comb. nov.Very large hypoconulids on M1 and M2, whereasthey are weakly developed in W. mcgrewi comb.nov. and W. shifrae comb. nov. M2 trigonid and tal-onid of similar width, whereas W. mcgrewi comb.nov. and W. shifrae comb. nov. have significantlywider M2 talonid bases relative to the trigonid base.M3 with larger paraconid and taller metaconid rela-tive to the protoconid than in W. shifrae comb. nov.Hypoconulid lobe of M3 not expanded as far dis-tally from the apex of the hypoconulid as in W.shifrae comb. nov. Differs from W. mcgrewi comb.nov. (but not W. shifrae comb. nov.) in havingupper molars with smaller distolingual basins. Inmesial view, M1 with a less clearly delineated pre-protocrista, like W. mcgrewi comb. nov. M2 with amuch more lingually expanded protocone lobe than

FIGURE 4. Environmental SEM images of four teeth ofWalshina esmaraldensis gen. et sp. nov. 1: left M3,SDSNH 76276; 2: right M1, SDSNH 76337; 3: right M2,

SDSNH 76338; 4: right M3, SDSNH 72583. All teeth are

in occlusal view. Arrow indicates the location of thefovea.

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in W. shifrae comb. nov. (but not in W. mcgrewicomb. nov.).

DESCRIPTION AND COMPARISONS

The only known M1 of W. esmaraldensis gen.et sp. nov. comes from LACM (CIT) 180 from SimiValley, Ventura County, California, and wasdescribed by Mason (1990). It differs from all other

FIGURE 5. Photographs taken with digital camera (1, 4-11) and Micro-CT scan reconstructions generated using Avizo7 (2, 3). Walshina shifrae comb. nov. (1, 4, 7, 10) – 1: right M1, CM 15797 (holotype; mirrored), in occlusal view; 4: leftM2, CM 15103, in occlusal view; 7: left M2, CM 21637, in occlusal view; 10: left M3; CM 15726, in occlusal view.

Walshina esmaraldensis gen. et sp. nov. (2, 5) – 2: left M1, LACM 40198 (holotype), in occlusal view; 5: left M2,SDSNH 62850, in occlusal view. Walshina mcgrewi comb. nov. (3, 6, 8, 9, 11) – 3: left M1, CM 15635 (holotype), inocclusal view; 6: left M2, CM15794, in occlusal view; 8, 9, 11: left mandibular fragment with M2, CM 29005, in occlusal

(8), buccal (9), and lingual (11) views.

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upper molars of W. esmaraldensis gen. et sp. nov.in having a less expanded distolingual basin, and ametastylar lobe that extends buccally to a similardegree as the parastylar lobe, which is a typical dif-ference between M1s and M2s in many early pri-mate taxa. Mason (1990) attributed LACM 40198to Ph. sp. cf. Ph. shifrae, and stated that the solediagnostic character to differentiate W. mcgrewicomb. nov. from W. shifrae comb. nov. was size.Although size is a useful diagnostic character inWalshina gen. nov., there are a number of otherdistinguishing features (i.e., position of the hypoco-nulid of M2, size of the hypoconulid on M1-2, trigo-

nid-talonid width proportion on M2, size ofparaconid on M3, height of metaconid on M3, sizeof hypoconulid lobe on M3, expansion of distolin-gual basins on upper molars, expression of thepreprotocrista on M1, expansion of the protoconelobe on M2) that can be used to differentiateamong the species. Nonetheless, Mason (1990)provides an adequate description of this tooth.

Both specimens of M2 have an oblique buccalborder due to a reduced metacone relative to theparacone and a more expanded distolingual basinthan M1. On the M2, the postprotocingulum runs

FIGURE 6. Micro-CT scan reconstructions of specimens of Walshina esmaraldensis gen. et sp. nov. generated usingAvizo 7: 1: left M1, LACM 40198 (holotype), in occlusal view; 2: left M2, SDSNH 87336, in occlusal view; 3: lingualfragment of a left M2, SDSNH 87337, in occlusal view; 4: lingual fragment of a right M2, SDSNH 42268, in occlusalview; 5, 6, 9, 10: left M2, SDSNH 87332, in occlusal (5), buccal (6), mesial (9) and lingual (10) views; 7, 8, 11, 12:

mesial fragment of a left M1, SDSNH 87331, in occlusal (7), buccal (8), mesial (11) and lingual (12) views; 13, 14, 17,

18: left M3, SDSNH 87334, in buccal (13), occlusal (14), lingual (17) and mesial (18) views; 15, 16, 19: distal fragment

of a right M3, SDSNH 87335, in buccal (15), occlusal (16) and lingual (19) views.

9


straight mesiodistally, creating a sharper angle withthe postcingulum than seen on the M1, where thepostprotocingulum and postcingulum are morebuccally oblique. The outline of M2 is rectangularrather than squared, with a longer buccolingual dis-tance. The distolingual basin is deep, with astraight distal aspect of the tooth, instead of con-cave. The paracone and metacone are well sepa-rated. The trigon basin is also deep. The parastylarregion is damaged in all M2 specimens, so it isimpossible to determine the presence of a para-style or the true degree of expansion of the para-stylar region. There is no metastyle. Only oneconule is present, a minute paraconule, with anassociated preparaconule crista, but no post-paraconule crista. The paraconule is closer to theparacone than to the protocone. The protocone isslightly skewed mesiolingually. The protocone lobeof W. esmaraldensis gen. et sp. nov. is very lin-gually expanded; more so than in W. shifrae comb.nov. but less so than in Trogolemur. However, noneof the other trogolemurins have a distolingual basinas expanded as that of the M2 of W. mcgrewicomb. nov., which is so expanded that the postcin-gulum curves distally beyond the plane of themetacone in the lingual aspect of the tooth. In con-trast, the distolingual basin in W. esmaraldensisgen. et sp. nov. does not extend further distallythan the metacone.

The only M3 known for the genus Walshinagen. nov. belongs to W. esmaraldensis gen. et sp.nov. The paracone and metacone of this tooth(SDSNH 76267) are very worn. The metacone issignificantly smaller than the paracone. No conulesare present. There is a strong postprotocrista, butthe preprotocrista is completely absent. The lingualhalf of the M3 is markedly shorter mesiodistallythan the buccal half, so that the tooth has a quitetriangular outline.

On the M1, the breadth of the talonid near theapices of the cusps is greater than the breadth ofthe trigonid. The paraconid is present on a well-developed paracristid that extends well beyond theparaconid and metaconid, making the front of thetooth very long and narrow. On the M1, the parac-ristid can be as tall as the tip of the paraconid, mak-ing that cusp poorly defined; whereas on the M2,the paracristid is lower and the paraconid is moredistinct from this crest. On the M1, the metaconid isdistally displaced relative to the protoconid and thepostvallid is stepped (sensu Silcox, 2001). Thecrests on the lower molars are all well delineated,in association with the teeth being fairly high

crowned (much more so, for example, than inSphacorhysis). On both M1 and M2, the distalslopes of the entoconid and the hypoconid areexpanded beyond the apices of these cusps, givingthe distal border of the tooth a convex aspect.There is a fovea on the distal aspect of the toothbetween the hypoconulid and the hypoconid. Thehypoconulid is well defined and shifted far buccally,being close to the hypoconid.

The M3 paraconid is large. The trigonid has asteep distal aspect of the metaconid. The hypoco-nulid is enlarged into a lobe. The M3 is narrower atthe hypoconulid than at the level of the tip of thehypoconid and the tip of the entoconid (i.e., lowhypoconulid lobe/mid tooth ratio). The only otherknown M3 of Walshina gen. nov. belongs to W.shifrae comb. nov. Whereas there is a clear differ-ence in size, the two species also differ in the rela-tive proportions of the hypoconulid lobe. Thebuccolingual width of the hypoconulid lobe in W.shifrae comb. nov. is smaller than in W. esmar-aldensis gen. et sp. nov. relative to the distancebetween the hypoconid and the entoconid, makingthis part of the tooth appear narrower. The hypoco-nulid lobe of W. shifrae comb. nov. also has a lon-ger distal slope than in W. esmaraldensis gen. etsp. nov., so that the hypoconulid lobe extends fur-ther from the tip of the hypoconulid.

Body mass estimates using equations by Gin-gerich et al. (1982) and the prosimian equation byConroy (1987) are shown for all species of trogole-murins in Table 1. Measurements used for calculat-ing body mass estimates are shown in Appendix 1.The equation by Conroy (1987) would be expectedto provide more relevant estimates, because it isbased on the subset of living primates most likelyto be comparable to the fossil taxa. However, heonly provided an equation for the M1, which isproblematic in this case because that tooth locus isunknown in several species of trogolemurins. Inparticular, although the smallest estimated bodymass is 58 g for Tr. myodes using the equation byConroy (1987), if an M1 was known for W. shifraecomb. nov., its estimated body mass would cer-tainly be lower. This is because the estimatesbased on teeth known for both species using theGingerich et al. (1982) equations are consistentlymuch lower for W. shifrae comb. nov. than for Tr.myodes (e.g., estimate based on M2 is 142 g for Tr.myodes compared to 109 g for W. shifrae comb.nov.). This would make W. shifrae comb. nov. thesmallest of the trogolemurins, and smaller than allother North American omomyoids (based on com-

10


parison to body mass estimates in Jones et al.,2014).

PHYLOGENETIC RELATIONSHIPS OF WALSHINA GEN. NOV.

In order to assess the phylogenetic position ofWalshina gen. nov. among Omomyoidea, we con-ducted a cladistic analysis based on the study byNi et al. (2004), as modified by Holroyd and Strait(2008), who revised the coding for two species(Purgatorius janisae and An. savagei), and addednew data for species of Loveina. We chose thisparticular matrix rather than more recent studies(e.g., Ni et al., 2013, 2016) because it includes atrogolemurin, Tr. myodes. Having coding from amember of this family available in the originalmatrix was important to ensure that we wereassessing characters in the same way as the origi-nal authors. The analysis also includes representa-tive tarsioids, anthropoids, adapoids, crownstrepsirrhines, and plesiadapiforms. Scandentiawas used as an outgroup to Primates. The rest ofthe trogolemurin primates were added to the matrixof Holroyd and Strait (2008): Tr. amplior, Tr. fragilis,Tr. leonardi, Sphacorhysis burntforkensis, W.esmaraldensis gen. et sp. nov., W. mcgrewi comb.nov., and W. shifrae comb. nov. The microchoerineMelaneremia bryanti (Hooker, 2007), and the anap-tomorphins An. sublettensis (Gazin, 1952, 1958),An. wortmani (Bown and Rose, 1984), and An.natronensis (Beard et al., 1992) were added aswell. The coding for Tr. myodes was partially reas-sessed, as was the one for Nannopithex sp. (toinclude generic variation seen in Na. zuccolae[Hooker, 2007]), and Scandentia was re-coded forcharacter 13. A total of 303 characters (194 dental,49 craniomandibular, 56 postcranial, and 4 soft tis-

sue and physiological [only coded for extant taxa]characters) were scored for a total of 66 taxa(matrix available on publication from www.morpho-bank.org under project number P3264).

A parsimony analysis was performed usingTNT v. 1.5 (Goboloff and Catalano, 2016) with allcharacters equally weighted and ordered, followingthe original analysis (Ni et al., 2004; Holroyd andStrait, 2008). Multiple character states were set tobe interpreted as polymorphisms, instead of uncer-tainties. A traditional search was implemented with1000 repetitions. The tree-bisection-reconnection(TBR) algorithm was used for branch swapping,with 1000 trees to save per replication.

Results of the Phylogenetic Analysis

The cladistic analysis yielded 90 equally parsi-monious trees. A strict consensus tree was gener-ated in TNT v. 1.5 from these trees (Figure 7, seealso Appendix 2). In the strict consensus tree,Walshina gen. nov. is found to be monophyletic,although the relationships among species ofWalshina gen. nov. remain unclear. The tribe Tro-golemurini was also recovered as monophyletic.

The inclusion of the rest of trogolemurins andthe four additional species of omomyoids, alongwith the revisions made by Holroyd and Strait(2008) and in the present paper, caused a fewchanges to the tree with respect to the topologyfound by Ni et al. (2004). The position of Purgato-rius is resolved in our tree, being the sister group tothe rest of Primates sensu lato. The rest of Plesi-adapiformes (excluding Purgatorius) form oneclade, sister group of Altanius orlovi, as in Ni et al.(2004). However, phylogenetic analyses of plesi-adapiforms with greater taxon sampling for thatgroup support the interpretation that this group ofprimates is paraphyletic and excludes An. orlovi,

TABLE 1. Estimated body masses (in g) of all species of Trogolemurini.

Gingerich et al.’s (1982) equations Conroy’s (1987)

M1 equation M1 M2 M3 M1 M2

Walshina esmaraldensis gen. sp. nov.

154 197 254 117 230 72

Walshina mcgrewi comb. nov. 237 275 - 245 315 115

Walshina shifrae comb. nov. - 109 199 65 108 -

Trogolemur myodes 125 142 263 125 231 58

Trogolemur amplior - 290 - - - -

Trogolemur fragilis - - 227 - - -

Trogolemur leonardi - - - 278 - -

Sphacorhysis burntforkensis 166 270 268 - - 78

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12

FIGURE 7. Hypothesis of relationships of Walshina gen. nov. in the context of the Order Primates. Strict consensuscladogram based on data modified from Holroyd and Strait (2008), including the addition of seven newly-coded tro-golemurins (Trogolemur amplior, Tr. fragilis, Tr. leonardi, Sphacorhysis burntforkensis, Walshina esmaraldensis gen.et sp. nov., W. mcgrewi comb. nov., and W. shifrae comb. nov.), a microchoerine (Melaneremia bryanti), and threeanaptomorphins (Anemorhysis sublettensis, An. wortmani, and An. natronensis). Trogolemurins are marked in blue.


which is interpreted as a euprimate (Silcox, 2001;Bloch and Boyer, 2002; Bloch et al., 2007, Silcox etal., 2010; Chester and Bloch, 2013; Chester et al.,2015; Ni et al. 2016). The tree differs from theresults obtained by Ni et al. (2004) in resolving theposition of Donrussellia as the sister taxon to therest of Euprimates (except for Altanius). Our treealso resolves Mahgarita and Rooneyia as sistertaxa. Although Rooneyia has been considered anomomyoid, it appears as being part of the strepsir-rhine radiation, as it did in Ni et al. (2004). All spe-cies of Teilhardina appear as basal haplorrhines,with unclear relationships among them. Eosimiasappears in our tree as the sister taxon of crownhaplorrhines (=Anthropoidea + Tarsius). Washaki-ins are monophyletic, although Loveina was foundto be paraphyletic (consistent with the results ofHolroyd and Strait, 2008). Our tree also resolves aclade that includes omomyins and macrotarsiins, inwhich omomyins are paraphyletic. Washakiins areresolved as the sister group to a subgroup of anap-tomorphins, which include Uintanius, Tetonius,Strigorhysis, Aycrossia, Anaptomorphus, andAbsarokius. The rest of anaptomorphins (Anemo-rhysis, Tetonoides, and Arapahovius) form anotherclade that is the sister group of trogolemurins.However, Anemorhysis does not appear to bemonophyletic. The sister group to the clade (Trogo-lemurini + Anemorhysis + Tetonoides + Arapa-hovius) is the Microchoerinae from Europe. Theposition of trogolemurins is, therefore, in contrastwith their position in the tree found by Ni et al.(2004), in which Trogolemur clustered togetherwith the European microchoerines. Finding trogole-murins to be more closely related to North Ameri-can anaptomorphins, rather than to Europeanmicrochoerids is more biogeographically parsimo-nious, reducing the number of implied dispersalevents between North America and Europe.

DISCUSSION

Trogolemurins have many dental features thatconverge on those of paromomyids, resulting intaxonomic confusion in the related literature: largehypoconulid lobes on M3, procumbent lower inci-sors, upper molars with a well-developed postpro-tocingulum and a fairly expanded distolingualbasin. The most diagnostic tooth for paromomyidsis the P4, which is enlarged with a tall, pointed,upright protoconid, and is very different from a tro-golemurin P4, which is proportionally smaller with amesially inclined trigonid. However, trogolemurinmaterial has classically been scarce and incom-

plete, often missing the diagnostic P4 (Robinson,1968; Krishtalka, 1978; Mason, 1990; Storer, 1990;Walsh, 1991a; Beard et al., 1992) or the upperdentition (Matthew, 1909; Beard et al., 1992), andin many cases taxa are known only from unassoci-ated isolated teeth (Robinson, 1968; Krishtalka,1978; Mason, 1990; Storer, 1990; Walsh, 1991a).Trogolemurin P4s are only known for Trogolemurand Sphacorhysis (Matthew, 1909; Emry, 1990;Gunnell, 1995), but not for Walshina gen. nov. Itwas not until work by Emry (1990) that the firstupper molars ascribed to a trogolemurin weredescribed, and not until work by Gunnell (1995)that the first upper dentition of Trogolemur in atooth row (including the first P4 known for a trogole-murin) was described. This lack of knowledgeabout the form of the upper molars in trogolemurinsled Storer (1990) to describe two complete andother fragmentary isolated upper molars as a newspecies of paromomyid, Ph. leonardi, from the latemiddle Eocene (Duchesnean) of Saskatchewan.Beard et al. (1992) transferred Ph. leonardi to theeuprimate genus Trogolemur and were the firstauthors to recognize the similarities between thesetwo groups.

It is now clear that in addition to the differ-ences in the morphology of the P4, there areimportant differences in upper and lower molarmorphology. Paromomyids have a more quadran-gular outline of the upper molars. Although bothparomomyids and trogolemurins have expandeddistolingual basins, the degree of expansion in tro-golemurins is most comparable to the oldest paro-momyids with plesiomorphic morphologies (i.e.,Paromomys), which are millions of years older thanthe oldest trogolemurins: the oldest paromomyidsare from the earliest Torrejonian, approximately 63Ma (Clemens and Wilson, 2009) whereas the old-est trogolemurins are from the latest Wasatchian,approximately 51 Ma (Williams and Covert, 1994).Trogolemurins are further characterized by havinga large lingual expansion of the protocone lobe onthe upper molars, which is not seen in any paromo-myids.

The M1 of trogolemurins differs from that toothin paromomyids in having a very oblique, steppedpostvallid, produced by a metaconid that is locatedwell distal to the protoconid. The M1 also has avery tall, strong paracristid, which is never soexpanded in paromomyids. The hypoconulid on theM1 and M2 of trogolemurins is also better devel-oped and more distinct, whereas paromomyidsshow weak to absent hypoconulids on these teeth.

13


The talonid of the M2 is shorter in trogolemurinsrelative to the length of the trigonid, making the tri-gonid and talonid closer in length in trogolemurins.

In sum, it is clear that in spite of the similaritiesbetween trogolemurins and paromomyids, thereare numerous characters that distinguish themfrom one another. An additional element to theargument that the material previously ascribed to I.mcgrewi and Ph. shifrae does not belong in Paro-momyidae is that it demonstrates morphology thatwould not be expected in paromomyids from solate in the family’s evolutionary history. In particu-lar, paromomyids generally exhibit increases in thedegree of expansion of the upper molar distolin-gual basin through time, and yet the upper molarsknown for the Uintan and Duchesnean purportedparomomyids are most comparable to those of theearliest paromomyids.

Re-classifying I. mcgrewi and Ph. shifrae asmembers of the new trogolemurin genus Walshinagen. nov. reframes our understanding of the timingof the demise of paromomyid diversity. Ignaciusmcgrewi and Ph. shifrae were the only members ofthe family identified from the Uintan and the Duch-esnean, so transferring them to Omomyoidea elim-inates the entire Uintan and Duchesnean record ofParomomyidae. It is worth noting, however, thateven when they were considered paromomyids,there was a marked discontinuity in the record ofthe family between the supposed Uintan forms andthe next oldest material. The first record of a paro-momyid comes from the Torrejonian of Montana,specifically the Torrejonian 1 biozone (Clemensand Wilson, 2009), and from that point there is acontinuous presence of this family until the latestWasatchian (Silcox et al., 2008). Then, the nextoccurrence of a paromomyid is in the middle Brid-gerian (Br2; Gunnell et al., 2009). The absence ofparomomyids from the very large samples of mam-mal specimens from later in the Bridgerian (Mat-thew, 1909; West, 1976; Emry, 1990; Gunnell andBartels, 1999; Gunnell et al., 2009) made theapparent reappearance of the group in the late Uin-tan somewhat surprising. Re-classifying the Uintanand Duchesnean material as Trogolemurini makesit clear that paromomyids were much reduced inrichness after the early Bridgerian in North Amer-ica. The only remaining post-Bridgerian record of aparomomyid is an isolated P4 from the Chadronianof North Dakota, tentatively ascribed by Kihm andTornow (2014) to the genus Ignacius, which is sep-arated from the next youngest specimen by someof the Bridgerian, and the entire Uintan and Duch-esnean, indicating a very long ghost lineage that

extends over a period of approximately 12 millionyears (middle Eocene boundary at 49 Ma, Clyde etal., 2001; Duchesnean-Chadronian boundary at 37Ma, Prothero and Emry, 2004). The North Dakotanspecimen shows clear features that ties it to paro-momyids, and in particular to the genus Ignacius.The outline of the tooth is squared, instead of rect-angular (as seen in Trogolemur, Gunnell, 1995),the cusps and styles are blunter, the parastylarlobe is very broad, and the postparacone andpremetacone cristae meet obliquely, making a Vshape in occlusal view, a trait observed in Ignacius.

Regardless of the apparent existence of thisone surviving lineage of geographically isolatedparomomyids, this family of plesiadapiforms fol-lows a clear trend of declining biodiversity after theend of the early Eocene in North America (lateWasatchian-early Bridgerian), which is clarified bythe re-classification of the Uintan and Duchesneanmaterial. Paromomyids reached their highest peakof biodiversity during the Wasatchian 4 (Wa4) bio-zone, whereas only one species is found in themiddle Bridgerian, Elwynella oreas (Rose andBown, 1982; Gunnell et al., 2009). The drop in bio-diversity of paromomyids could therefore be cor-related with the Early Eocene Climatic Optimum(EECO), the warmest sustained period of the entireCenozoic (Clyde et al., 2001; Zachos et al., 2008;Hyland and Sheldon, 2013; Chew, 2015). Paromo-myids are also extremely rare during the Paleo-cene-Eocene Thermal Maximum, only known froma lower molar of I. graybullianus and an uppermolar and an edentulous jaw of Ph. praecox (Gin-gerich, 1989; Rose et al., 2012). The drop in paro-momyid biodiversity during the Wasatchian 5biozone also coincides with the warming periodthat precedes the EECO (including at least onehyperthermal; Chew and Oheim, 2013). In con-trast, the highest peak of paromomyid biodiversity(Wa4) occurs during a cooling period between Bio-horizon A and Biohorizon B (Chew and Oheim,2013). And although the material has yet to bedescribed in detail, paromomyids are also the onlyprimates known from the early-middle Eocene ofEllesmere Island in the Canadian High Arctic (Westand Dawson, 1977; McKenna, 1980). That part ofthe world is interpreted to have been much milderin the Eocene than today, but winter temperaturesare still inferred to have been near freezing (Eberleand Greenwood, 2012), making this a chilly envi-ronment for primates. These various lines of evi-dence suggest that, as a family, paromomyidspreferred cooler temperatures, and that the near

14


extinction of the family in North America after theWasatchian-Bridgerian boundary could potentiallybe ascribed to increasing global temperatures.

Late Paromomyids from Europe

Although the reinterpretation of most of thelatest occurring purported paromomyid materialfrom North America makes it clear that the familywas largely extinct on that continent after the earlyBridgerian, there are some later occurring speci-mens from Europe that have been attributed to thefamily, which could signal a longer persistence onthat continent. The only known genus of paromo-myid from Europe is Arcius (Russell et al., 1967;Godinot, 1984; Estravís, 2000; Marigó et al., 2012).Interestingly, the occurrence of paromomyid spe-cies in Europe through time shows some similari-ties to what has been observed in North America.Arcius first appears in Europe during the earliestEocene (Mammal Paleogene zone 7 [MP7], Neus-trian: Estravís, 2000), followed by a continuouspresence of this family until the Grauvian (MP10;early-middle Eocene; Godinot, 2015). After that, noparomomyids are found during MP11 or MP12. It isnot until MP13 (Geiseltalian) that a few teeth arereported to belong to Paromomyidae (Sudre, 1978;Rémy et al., 1997). A second lower molar of Arciussp. from Saint-Maximin (France) is illustrated andreported in a faunal list by Rémy et al. (1997), andtwo third lower molars (one of them illustrated inAumont, 2003) from Bouxwiller (France) appear ina faunal list in Sudre (1978). However, this scarcematerial, comprised of three isolated molars fromthe middle Eocene of Europe, might not be paro-momyids.

The tooth from Saint-Maximin, identified hereas an M2 (and as either an M1 or an M2 by Rémy etal., 1997), is strikingly different from that tooth inany Arcius species. It has a prominent paraconid,which is not consistent with what it is observed in atypical Arcius M2. Also, the trigonid area is signifi-cantly larger in relation to the talonid than in anyspecies of Arcius. We suggest that it could be adiminutive adapoid, and tentatively ascribe it to cf.Anchomomys sp., based on the presence of thisgenus at that site. The M3s from Bouxwiller wereidentified as Arcius lapparenti by Aumont (2003).However, they are very flat and significantly shortermesiodistally than those of Ar. lapparenti. The tri-gonid is shorter mesiodistally in relation to its widththan in Ar. lapparenti, and the buccolingual dimen-sion from the hypoconid to the entoconid is signifi-cantly greater than in any species of Arcius. Also,the hypoconulid lobe, although expanded, is signifi-

cantly smaller than the ones seen in any Eoceneparomomyids. Therefore, we suggest that it couldpotentially be a microchoerid omomyoid, and tenta-tively ascribe these teeth to cf. Nannopithex sp.,based on the presence of this genus at that site.The fact that the Geiseltalian material likelybelongs to diminutive euprimates rather than to theParomomyidae is consistent with the near extinc-tion of paromomyids after the early middle Eoceneobserved in North America.

CONCLUSIONS

The material from southern California previ-ously ascribed to Phenacolemur by Mason (1990)and Walsh (1991a) are not paromomyids. Theyrepresent a new species of trogolemurin omo-myoid, W. esmaraldensis gen. et sp. nov. The Cali-fornian sample of Walshina gen. nov. adds to ourknowledge of the variability and diversity of omo-myoids, particularly trogolemurins, and significantlychanges our understanding of the temporal distri-bution of the plesidapiform family Paromomyidae.The former paromomyids I. mcgrewi and Ph.shifrae from the Uintan and Duchesnean of Wyo-ming are transferred to the genus Walshina gen.nov., eliminating any record of paromomyids fromthe Uintan and the Duchesnean. The nearly com-plete lack of paromomyids after the onset of theBridgerian could be correlated with the warmestsustained period of the Cenozoic, the EECO. Thispattern is consistent with the disappearance ofparomomyids after the Grauvian in Europe, makingthis a generalized trend of extinction or near extinc-tion of this group worldwide in the middle Eocene.

Trogolemurin omomyoids and paromomyidplesiadapiforms converge in general dental mor-phology, suggesting that they might have been eat-ing similar foods. It is notable that omomyoids withthis morphology appear only after the near extinc-tion of paromomyids in North America, suggestingthat perhaps the marked decline in paromomyidsleft niche space available for omomyoids. The soleremaining late occurrence of a paromomyid, fromthe Chadronian of North Dakota, has been inter-preted as being part of a relictual fauna surviving inthe Great Plains, distinct from the Rocky Mountainprovince (Kihm and Tornow, 2014). Perhaps sur-vival of a paromomyid in that context was madepossible by its isolation from ecologically similartrogolemurin omomyoids.

Finally, our phylogenetic analysis shows thatWalshina gen. nov. is monophyletic, nested insideTrogolemurini. The most closely related group totrogolemurins is a clade of anaptomorphins com-

15


posed by Anemorhysis, Tetonoides, and Arapa-hovius. Therefore, our phylogenetic analysis isconsistent with the previously inferred close rela-tionship of trogolemurins to Anemorhysis (Wil-liams, 1994; Williams and Covert, 1994; Tornow,2008) and to Tetonoides and Arapahovius (Gun-nell, 1995). Although the most recent definition ofTrogolemurini by Gunnell and Rose (2002) is fairlyconstricted in number of genera, our phylogeneticresults are consistent with a broader definition ofTrogolemurini suggested by Beard et al. (1992), inwhich Anemorhysis, Tetonoides, and Arapahoviusmight also be included.

ACKNOWLEDGMENTS

We would like to thank W.A. Clemens(UCMP), T.A. Deméré, and K.A. Randall (SDSNH),T. Tokaryk, R.C. McKellar, and H. Bryant (RSM),A.R. Tabrum (CMNH), J.J. Eberle, T. Culver, and T.Karim (UCM) for access to specimens and/orcasts. We thank T.M. Bown, M.A. Tornow, and T.B.Viola for relevant discussions. Thanks to D.R.

Begun, W.A. Clemens, G.F. Gunnell, and M.A.Schillaci for comments on a previous version of thismanuscript. We are thankful to the Editorial Crewof Palaeontologia Electronica and three anony-mous reviewers for comments that substantiallyimproved this paper. Thanks to R. Bhagat for themaking of relevant casts. We are very grateful toJ.T. Gladman for his assistance in CT-scanning thespecimens of Walshina gen. nov. Thanks to K.R.Selig for further assistance during scanning. Photo-graphs were taken and edited by G. San MartinFlores, D. Lin, PAH, MTS, and SLT. We are gratefulto D.M. Boyer, G. Dewar, and J.A. Teichroeb foraccess to resources. Thanks to R. Blakey (Colo-rado Plateau Geosystems, Inc., USA) for the per-mission to use paleogeographical maps. Thisresearch was supported by the Doris O. and Sam-uel P. Welles Research Fund and a University ofToronto Department of Anthropology ResearchTravelling Grant to SLT, and an NSERC DiscoveryGrant to MTS.

REFERENCES

Abbott, P.L. 1999. Rise and fall of San Diego: 150 million years of history recorded in sedimentary rocks. 1st Edition. Sunbelt Publications, San Diego, California.

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APPENDIX 1.

APPENDIX TABLE 1. Measurements (in mm) for upper molars of Trogolemurini.

Specimen Species M1L M1W M2L M2W M3L M3W Locality Reference Notes

CM 14598 Walshina shifrae comb. nov.

- - 1.24 1.81 - - 5(A?), Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

Identified as Ignacius mcgrewi in Robinson (1986)


- - 1.40 2.00 - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)

CM 15635 Walshina mcgrewi comb. nov.

2.00 2.78 - - - - 5-front, Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

Holotype


- - 1.80 2.90 - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)


2.00 - - - - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)


2.00 - - - - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)


1.30 2.00 - - - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)

Holotype


- - 1.40 - - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)


1.20 1.90 - - - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)

LACM 40198 Walshina esmaraldensis gen. et sp. nov.

1.60 2.20 - - - - LACM (CIT) 180, north of Simi Valley, Ventura Co, CA, USA

This paper Holotype

SDNHM 42268

Walshina esmaraldensis gen. et sp. nov.

- - 1.66 - - - SDNHM loc. 3426, Collwood South, San Diego Co, CA, USA

This paper

SDNHM 62850


- - 1.64 2.39 - - SDNHM loc. 4020, State Route 125, San Diego Co, CA, USA

This paper

SDNHM 76267


- - - - 1.31 2.05 SDNHM loc. 4082 (Emerald Ridge 2), northwest San Diego Co, CA, USA

This paper

SDNHM 87336


- - 1.76 2.48 - - SDNHM loc. 4925, Kelly Ranch Core, San Diego Co, CA, USA

This paper

SDNHM 87337


- - 1.54 - - - SDNHM loc. 4925, Kelly Ranch Core, San Diego Co, CA, USA

This paper

SMNH P1899.1007

Trogolemur leonardi

1.90 3.17 - - - - SMNH loc. 72J04-0006, east side of Lac Pelletier, SK, Canada

Storer (1990)

SMNH P1899.1016

Trogolemur leonardi

- - - - 1.58 2.57 SMNH loc. 72J04-0006, east side of Lac Pelletier, SK, Canada

Storer (1990)

Identified as M2 in Storer (1990)

24


UCM 25049 Walshina shifrae comb. nov.

- - 1.24 1.67 - - 5A, Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

Identified as Ignacius mcgrewi in Robinson (1986)

USNM 417390

Trogolemur myodes

- - 1.60 2.60 - - Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

USNM 417391

Trogolemur myodes

- - - - 1.30 2.20 Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

Incorrectly listed as a lower m3 in Emry (1990)

USNM 417396

Trogolemur myodes

1.60 2.30 - - - - Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

Identified as M2 in Emry (1990)

Specimen Species M1L M1W M2L M2W M3L M3W Locality Reference Notes

25


A1

C1

C2

C4

C4

S7

S7

S7

S7

S7

S8

S8

APPENDIX TABLE 2. Measurements (in mm) for lower dentition of Trogolemurini.S

pe

cim

en

Sp

ec

ies

p3

L

p3

W

p4

L

p4

W

m1

L

m1

W

m2

L

m2

W

m3

L

m3

W

Lo

cali

ty

Re

fere

nc

e

No

tes

MNH 2599

Trogolemur myodes

1.05 0.78 1.33 1.14 1.66 1.23 1.60 1.44 2.07 1.27 Granger S 6, Uinta Co, WY, USA

This paper Holotype

M 5793

Walshina mcgrewi comb. nov.

- - - - - - 2.00 1.80 - - Badwater Creek, Natrona Co, WY, USA

Krishtalka (1978)

M 1637

Walshina shifrae comb. nov.


Krishtalka (1978)

M 0069

Trogolemur amplior

- - - - 2.20 1.85 1.95 1.95 - - CM loc. 34, Wind River Basin, Natrona Co, WY, USA

Beard et al. (1992)

M 1152

Trogolemur fragilis

- - - - - - - - 1.95 1.20 CM loc. 34, Wind River Basin, Natrona Co, WY, USA

Beard et al. (1992)

DNHM 2583


- - - - - - - - 1.80 1.18 SDNHM loc. 4081 (Emerald Ridge 1), northwest San Diego Co, CA, USA

This paper

DNHM 6266


- - - - - - - - 2.07 1.26 SDNHM loc. 4082 (Emerald Ridge 2), northwest San Diego Co, CA, USA

This paper

DNHM 6337


- - - - 1.83 1.48 - - - - SDNHM loc. 4082 (Emerald Ridge 2), northwest San Diego Co, CA, USA

This paper

DNHM 6338


- - - - - - 1.90 1.61 - - This paper

DNHM 6339


- - - - - - - - - 1.19 SDNHM loc. 4082 (Emerald Ridge 2), northwest San Diego Co, CA, USA

This paper

DNHM 7332


- - - - - - 1.79 1.46 - - SDNHM loc. 4925, Kelly Ranch Core, San Diego Co, CA, USA

This paper

DNHM 7333


- - - - - - 1.79 1.57 - - SDNHM loc. 4925, Kelly Ranch Core, San Diego Co, CA, USA

This paper

26


S8

U2

U2

U2

U2

U3

U4

U3

U4

U4

U4

Y1

DNHM 7334


- - - - - - - - 2.17 1.38 SDNHM loc. 4925, Kelly Ranch Core, San Diego Co, CA, USA

This paper

CM 5175


- - - - - - 1.50 1.33 - - 5A, Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

Identified as Ignacius mcgrewi inRobinson (1986)

CM 6012


- - - - 2.06 1.76 - - - - 5A, Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

CM 6432


- - - - - - 1.25 1.06 - - 5A, Badwater Creek, Natrona Co, WY, USA

Robinson (1968)

Identified as Ignacius mcgrewi inRobinson (1986)

CM 9005



Krishtalka (1978)

CM 8323



Krishtalka (1978)

CM 6602

Trogolemur amplior

1.20 0.95 - - - - - - - - UCM loc. 80062, Wind River Basin, Natrona Co, WY, USA

Beard et al. (1992)

M 0966

Sphacorhysis burntforkensis

- - 1.40 1.30 1.90 1.50 2.00 1.80 1.80 1.50 UM loc. BRW-264, sec. 22, T.13N., R.113W., Uinta Co, WY, USA

Gunnell (1995)

SNM 17355

Trogolemur myodes

1.00 0.80 1.20 1.20 1.70 1.50 1.40 1.60 2.00 1.30 Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

SNM 17356

Trogolemur myodes

1.00 0.90 1.10 1.25 1.50 1.50 1.40 1.60 2.10 1.40 Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

SNM 17389

Trogolemur myodes

- - - - 1.70 1.50 - - - - Elderberry Canyon Quarry, White Pine Co, NE, USA

Emry (1990)

PM VP 3523

Trogolemur myodes

- - - - - - 1.41 1.43 1.96 1.15 Sweetwater Co, WY, USA

This paper

Sp

eci

me

n

Sp

eci

es

p3

L

p3

W

p4

L

p4

W

m1

L

m1

W

m2

L

m2

W

m3

L

m3

W

Lo

ca

lity

Re

fere

nce

No

tes

27


APPENDIX 2.

Newick format for tree from Figure 7.

(Scandentia,(Purgatorius_spp,((Altanius_orlovi,(Plesiolestes_problematicus,('Plesiadapis/Pro-nothodectes',Chronolestes_simul))),(Donrussellia_sp.,(((Lemur_catta,(Galagoides_demidoff,Microcebus_murinus)),((Mahgarita_stevensi,Rooneyia_viejaensis),(Notharctus_sp.,(Pronycticebus_gaudryi,(Adapis_parisiensis,Leptadapis_magnus))))),(Teilhardina_americana,(Teilhardina_belgica,Teilhardina_asiatica),(Steinius_vespertinus,(Omomys_sp.,(Hemiacodon_gracilis,Macrotarsius_montanus))),(Eosimias_sp.,(Tarsius_sp.,(Proteopithecus_sylviae,(Catopithecus_browni,((Aegyptopithecus_zeuxis,(Apidium_phiomense,Parapithecus_grangeri)),(Dolichocebus_gaimanensis,(Callicebus_moloch,(Aotus_trivirgatus,Saimiri_sciureus)))))))),(((Uintanius_ameghini,(Tetonius_sp.,((Absarokius_sp.,Anaptomorphus_sp.),(Aycrossia_lovei,Strigorhysis_sp.)))),(Loveina_sheai,Loveina_minuta,Loveina_wapitiensis,(Dyseolemur_pacificus,Washakius_insignis,Loveina_zephyri,Shoshonius_cooperi))),((Nannopithex_sp.,Melaneremia_bryanti,Pseudoloris_parvulus,(Microchoerus_erinaceus,Necrolemur_antiquus)),((Anemorhysis_savagei,(Anemorhysis_wortmani,((Anemorhysis_sublettensis,Anemorhysis_natronensis),(Arapahovius_gazini,Tetonoides_sp.)))),(Trogolemur_myodes,Trogolemur_amplior,Trogolemur_fragilis,Trogolemur_leonardi,Sphacorhysis_burntforkensis,(Walshina_esmaraldensis,Walshina_mcgrewi,Walshina_shifrae)))))))))));

28

new omomyoids (euprimates, mammalia) from the late uintan

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